Factors determining the distribution of the green spruce aphid, Elatobium abietinum, on young and mature needles of spruce

1. The green spruce aphid Elatobium abietinum , was less active and more likely to feed when on mature rather than young needles of Sitka spruce.
2. Compared with survival on mature needles, survival of aphids on young needles was significantly reduced, especially on newly flushed needles, but became less so as the needles matured. The survival of aphids kept close to, but not in contact with, young needles was similarly reduced.
3. On young needles stripped of epicuticular wax, aphids were less active and fed and survived for longer than aphids on untreated young needles. In contrast, the supply of additional amino acids to shoots bearing young needles did not significantly improve the survival of aphids on these needles.
4. In a choice test, aphids were able to discriminate between epicuticular wax from young and mature needles. The reciprocal transfer of wax between young and mature needles reversed the aphids' marked preference for mature needles.
5. These results indicate that the aphid is deterred from settling and feeding on young needles by a volatile substance present in the epicuticular wax.     

Seasonal changes in the distribution of green spruce aphid Elatobium abietinum (Walker) (Homoptera: Aphididae) in the canopy of Sitka spruce

1 Seasonal changes in the distribution of green spruce aphid Elatobium abietinum (Walker) within the canopy of 20–25‐year‐old Sitka spruce are described based on data from two low‐altitude sites (310–420 m above sea‐level), two mid‐altitude sites (500–550 m a.s.l) and one high‐altitude site (610 m a.s.l). 2 Aphids were counted throughout the canopy on shoots representative of all needle age‐classes present at each whorl of branches. Counts were made during the middle week of each month from September to July for 4 years (1999–2003), and mean E. abietinum densities at each canopy position were calculated separately for each month and for the low‐, mid‐ and high‐altitude sites. 3 During September to November, the highest densities of E. abietinum occurred on 3–4‐year‐old needles on branches low in the canopy. Over the winter and spring, the centre of the aphid’s distribution shifted outward and upward, so that by June of the next year the highest aphid densities occurred on current and 1‐year‐old needles on branches near the top of the tree. 4 The aphid distribution was re‐set each year during July, at the time when the nutrient quality of the host was in decline and E. abietinum populations were decreasing. Aphid densities decreased less on 3–4‐year‐old needles than on current and 1‐year‐old needles, suggesting that older needles were a superior food resource at this time of year and in the autumn. However, other factors, such as higher temperatures in the upper canopy during the summer or differential mortality caused by natural enemies, could also have contributed to the change in distribution. 5 The outward and upward shift in the aphid distribution over the winter period provided no evidence that aphids at positions lower and deeper in the canopy were better insulated from freezing temperatures and had higher over‐winter survival rates. Mean air temperatures at the top and bottom of the canopy during the winter were also found to differ by only 0.1–0.2 °C. 6 The percentage of the total aphids per tree that occurred on current or 1‐year‐old needles varied widely between seasons and between sites. Consequently, sampling programmes designed to estimate total population numbers of E. abietinum have little option but to sample needles throughout the canopy, and at regular intervals during the period when the aphid is abundant.     

The effect of previously infested spruce needles on the growth of the green spruce aphid, Elatobium abietinum, and the effect of the aphid on the amino acid balance of the host plant

The effect of chlorosis induced in needles of Sitka and Norway spruce by the green spruce aphid on growth of the aphid is investigated, and the effect of infestation of the aphid on amino acid levels in Sitka spruce foliage is reported. On both Sitka and Norway spruce green spruce aphids were heavier when reared on chlorotic (previously infested) needles than when reared on green (previously uninfested) needles. The effect was more pronounced on Sitka than on Norway spruce. Infestation of the aphid altered the amino acid balance of Sitka spruce foliage but not the concentration of total amino acids. Possible causes of chlorosis, the influence of individual amino acids on aphid growth, the potential effect of chlorosis on outbreaks of the aphid and the differences in susceptibility of Sitka and Norway spruce to damage by the aphid are discussed.     

Effects of temperature on overwintering populations of the green spruce aphid Elatobium abietinum

Recorded minimum temperatures of – 7 ^oC or lower noticeably reduced overwintering populations of the green spruce aphid, Elatobium abietinum, in north-east Scotland. It is suggested that, at these temperatures, ice formation in the needles of the host Sitka spruce, caused attached aphids to freeze. Aphid mortality also occurred when maximum temperatures did not rise above + 6 ^oC for prolonged periods, possibly as a result of starvation following an extended chill coma. A diagrammatic representation of the main factors affecting anholocyclic populations of E. abietinum during the winter is presented to emphasize the governing role played by temperature. The balance between mortality and recruitment determines the size of the population at the end of the winter, and this in turn determines the subsequent summer infestation. It should be possible, therefore, to predict aphid outbreaks either from winter temperatures or from the number of aphids present at the end of the winter. Temperature records obtained from integrating thermometers indicated that the inside of the lower crown tended to be the warmest part of the tree during the winter, resulting in greater aphid survival in the lower branches.     

Relative suitability of crested wheatgrass and other perennial grass hosts for the Russian wheat aphid (Homoptera: Aphididae)

The Russian wheat aphid, Diuraphis noxia (Mordvilko) (Homoptera: Aphididae), reproduces parthenogenetically in North America and must survive year-round on host plants, including in late summer when small grains are not in cultivation. During this time, cool-season perennial wheatgrasses (Poaceae: Triticeae) contribute substantially to aphid survival, crested wheatgrass (Agropyron spp.) particularly. In greenhouse studies, the number of aphids per plant was measured after four infestation periods on unvernalized and vernalized wheatgrasses. Before placement on these test plant species, aphids were reared either on winter wheat or on the grass host species on which aphid progeny were counted. On vernalized plants, aphids reared on wheat resulted in more aphids per test plant than when the aphids were reared on wheatgrasses, but on unvernalized plants the number of aphids per test plant did not differ significantly regardless of rearing host. Aphids on crested wheatgrass were similar in number to the other grasses when plants were unvernalized. However, when plants were vernalized, crested wheatgrass supported significantly more aphids than some of the other hosts. Aphid numbers increased on all test species as infestation period lengthened, and plant growth was largely unaffected by aphid feeding. These results suggest if sufficient moisture is available during summer when small grains are not in cultivation, all host species observed are capable of sustaining aphids. Crested wheatgrass is an abundant and important host of the Russian wheat aphid in its northern range of the western United States, but other less prevalent wheatgrasses also may contribute to aphid survival during late summer when small grains are not in cultivation.     

A comparison of Elatobium abietinum populations on Sitka spruce trees differing in needle retention during aphid outbreaks

Summary No difference in the rate of symptom development and needle loss was evident between resistant and susceptible trees. Population growth rates, which were higher on current year needles in the upper crown, adult weights, reproductive state and instar ratios were similar for aphids from both resistant and susceptible trees. Overwintering survival rates were higher on susceptible trees and were the main cause of the higher number of aphids found on susceptible trees and on the lower crown in comparison with the resistant trees and the upper crown. The supercooling points of needles from resistant trees were higher than those from susceptible trees during the three coldest months of the year-December, January and February. As the aphid freezes at the same temperature as the needle on which it feeds differences in needle freezing temperatures can result in greater aphid mortality on the resistant trees at the temperatures prevailing in NE Scotland during the period of the study. No correlation was found between provenance latitude and supercooling ability of the needles although a very great variability was evident between individual trees. Changes in the form of the super-cooling temperature frequency curves indicated a gradual increase in the quantity of nucleators present in the spruce needles during the winter, the change being more pronounced in the resistant needles.     

Performance of the green spruce aphid, Elatobium abietinum (Walker) on previously defoliated Sitka spruce

Abstract 1 The green spruce aphid, Elatobium abietinum, is an important defoliator of Sitka spruce in the U.K. However, it is usual for years in which high E. abietinum populations occur to be followed by a year with low aphid densities. The possibility that the performance of E. abietinum is reduced on previously infested Sitka spruce, and that this is the cause of year‐to‐year fluctuations in population density, was investigated by comparing population development and the growth rate of individual aphids on experimentally defoliated trees. 2 Separate experiments were performed to determine whether aphid performance was reduced either in the autumn immediately after defoliation in the spring, or was reduced in the spring of the next year. Different rates of initial defoliation on trees used to test aphid performance were created by artificially infesting the trees with aphids in the spring before the experiments, and varying the time of infestation. 3 Population development and the mean relative growth rate (MRGR) of individual aphids on previously defoliated and undefoliated Sitka spruce did not differ significantly in the spring of the next year. No differences were observed in the nutrient content of the 1‐year‐old needles of previously defoliated or undefoliated trees at this time. 4 In the autumn and winter immediately after spring defoliation, aphid MRGR was significantly higher on trees that had been heavily defoliated earlier in the season compared with trees that had been lightly defoliated. However, the difference in MRGR decreased over the winter period. Nitrogen, phosphorous and potassium concentrations were 9.4–12.2% higher, at the beginning of the autumn, in the current year needles of heavily defoliated trees than in the current year needles of lightly defoliated trees. 5 The experiments indicate that high populations of E. abietinum in the spring do not induce any defensive mechanisms in Sitka spruce that adversely affect subsequent generations of the aphid. By contrast, the results suggest that high spring densities of the aphid improve the nutritional quality of the current year's foliage for autumn generations.     

Damage caused by the Green Spruce Aphid to Norway and Sitka spruce needles

Three-month-old needles of Sitka spruce were less susceptible to Elatobium abietinutn than 15-month-old needles. Symptoms appeared after longer aphid feeding times but only a proportion of damaged needles fell. After short feeding periods symptoms appeared in more Norway spruce needles than in Sitka spruce, whereas longer feeding periods resulted in more needles producing symptoms in the Sitka spruce. The symptoms took 4—6 days longer to appear in Norway spruce, and needle fall followed a longer feeding period than on Sitka spruce. Following 72 h feeding, needle fall occurred more quickly on Sitka spruce than on Norway spruce. The time taken for needle fall to occur was inversely related to the feeding time in Sitka spruce but such a response was not evident in Norway spruce. The results are discussed in relation to the differences exhibited in the probing behaviour of the aphid on the two spruces.     

Supercooling and the low-temperature survival of the green spruce aphid Elatobium abietinum

Laboratory investigations into the low-temperature tolerance of the green spruce aphid, Elatobium abietinum, revealed that the insect was killed by freezing. Aphids and host Sitka spruce needles showed similar seasonal changes in supercooling ability. A noticeable increase in this ability occurred between June and October. Aphids were more susceptible to low temperatures when attached to the plant. It is suggested that mortality resulted from ice which formed in the sap of the host needles and spread into the feeding aphids via their mouthparts. Neither the chlorotic banding of needles, caused by aphid feeding, nor needle length affected needle supercooling. Increased duration of exposure increased the probability of freezing of supercooled needles at low temperatures. A small percentage of first-instar nymphs supercooled to much lower temperatures than the remainder of the population. These were newly born nymphs whose high supercooling ability markedly decreased when they began to feed.     

The growth and decline of a population of the spruce aphid Elatobium abietinum during a three year study,and the changing pattern of fecundity,recruitment and alary polymorphism in a Northern Ireland Forest

The population size and structure of the green spruce aphid was followed throughout the spring — summer cycle on the same group of trees in a low-elevation coastal Sitka spruce forest for three consecutive years. The relationship between the pattern of change and the phenology of bud burst, which heralds a marked change in needle sap quality, suggests that yearly differences in the winter temperature regime may affect the duration of the population growth phase and hence the peak numbers attained in late spring.An index of population growth rate was sufficiently sensitive to aphid fecundity during the population cycles of two years to suggest that the changing rate of fertility was the decisive process governing changes in population size. The commitment of aphids to alate development was greater than that recorded elsewhere in Britain but did little to effect population decline since the contribution of alatae to larviposition was substantial while seasonally pulsed.     

The effect of needle age on the acceptability of Sitka spruce needles to the aphid,Elatobium abietinum (Walker)

Summary A comparison of the feeding behaviour of E. abietinum on current year needles and previous year needles of P. sitchensis revealed that during the summer months the aphids preferentially settled on previous year needles, this response not being evident in winter. Aphids on current year needles in summer took a much longer period of time to commence sap uptake than in winter, intake ceasing following a very short feeding period. On previous year needles sap uptake in December commenced after a shorter feeding period than in June. Analysis of total and soluble nitrogen levels in Sitka spruce needles showed that current year needles had initially higher levels during shoot elongation in May and early June, but that previous year needles had higher levels for most of the remainder of the year. Quantitative analyses of amino acids revealed that in current year needles the levels were generally lower than in previous year needles. Less marked proportional differences were observed between previous year needles in May and in July/August when the needles were unsuitable. Addition of amino acids in solution into cut current year shoots resulted in increased longevity on shoots containing introduced iso-leucine, histidine and methionine and revealed a general imbalance of the amino acids.     

Effect of exposure to fluoride,nitrogen compounds and SO2 on the numbers of spruce shoot aphids on Norway spruce seedlings

Summary Development of spruce shoot aphid (Cinara pilicornis Hartig) populations was monitored in natural and artificial infestations of Norway spruce (Picea abies Karst.) seedlings, exposed to air pollutants in an experimental field. The pollutants, applied both singly and in mixtures, were gaseous sulphur dioxide, NaF (30 mg l^-1 F) and Ca(NO₃)₂ or (NH₄)₂SO₄ in aqueous solutions (200 mg l^-1 N). Aphid numbers on 10 seedlings in each treatment and two control plots were counted at 2-week intervals. At the beginning of the experiment aphid numbers did not differ between treatments. Aphid populations peaked in late June and early July. All the pollutants and their combinations significantly increased the numbers of aphids per seedling. Four apterous females were transferred to spruce seedlings which were growing in containers in the same plots. After 4–5 weeks aphid numbers were significantly higher in the fluoride treatment and in the combined treatment of fluoride, nitrogen and SO₂. The pollution treatments did not have a significant effect on shoot growth. Concentrations of F and S in needles were higher in treatments involving these pollutants. There were no significant differences in the concentrations of free amino acids in shoot stems between control and fluoride treatment. However, the relatively low concentration of arginine in the F treatment at the end of the growing season might indicate disturbances in the nitrogen metabolism of spruce seedlings.     

Effects of Elatobium abietinum on nutrient fluxes in Sitka spruce canopies receiving elevated nitrogen and sulphur deposition

1 In a field acid mist simulation experiment, Sitka spruce (Picea sitchensis) was sprayed with different pollutant treatments: N, NH₄NO₃; S, Na₂SO₄; NS Acid, NH₄NO₃ + H₂SO₄ and control, no spray. Treatment effects on the abundance of the green spruce aphid Elatobium abietinum and honeydew production were assessed. In addition, needles were sampled for phyllosphere micro‐organisms. In a manipulative experiment, shoots were established and maintained as with or without E. abietinum infestation in order to determine the effects of infestation on needle loss and throughfall nutrient fluxes. 2 Aphid numbers were highest during the end of May and early June, with almost twice as many needles infested in the NS Acid treatment compared with the other treatments. Honeydew production was not affected by the treatments. 3 On infested shoots, increasing numbers of yellowing and dead needles were recorded above the throughfall collectors as the season progressed. The numbers of dead needles falling into the collectors were significantly higher beneath infested shoots. There were strong positive correlations between aphid numbers above the throughfall collectors, the number of yellowing and dead needles on the shoots and the number of needles in the funnels of the throughfall collectors. Litter production was more affected by aphid number than by pollutant treatment. 4 Bacteria, yeasts and filamentous fungi were more prolific on infested needles and treatment effects on colony forming units (CFUs) were most pronounced in the NS Acid treatment. 5 Fluxes of inorganic nitrogen beneath infested shoots were generally lower than beneath uninfested shoots. This effect was more pronounced in those treatments that supplied N i.e. N, NS Acid. The combination of aphid infestation and N‐addition exerted the strongest influence on nutrient fluxes. The fluxes of potassium and of organic carbon (DOC) were higher beneath infested shoots in all treatments, through most of the survey period.     

Aphid honeydew and its effect on the phyllosphere microflora of Picea abies (L.) Karst

Aphids of the genus Cinara, feeding on Norway spruce, excrete copious amounts of honeydew, a carbon-rich waste product, which accumulates locally on needles and twigs. We investigated the role of honeydew as a potential source of energy which might promote the growth of micro-organisms in the phyllosphere of conifer trees. To approach this question, we followed the population dynamics of Cinara spp. in a natural forest stand over two seasons. We also studied the amounts of honeydew produced by individual aphids and identified potential parameters which might influence honeydew production. Finally, we determined the growth of micro-organisms on infested and uninfested needles of Norway spruce during the growing season. Confined to Picea abies, the investigated Cinara species only became abundant in midsummer, when needles and shoots were expanding. The populations showed only a single peak in abundance, the timing and magnitude of which may vary from year to year due to weather conditions, changes in plant quality in a yearly cycle or the impact of natural enemies. The amount of honeydew produced by individual aphids was dependent on the developmental stage of the aphid, the nutritional supply of its host plant and on the developmental state of the Norway spruce (e.g. bud burst, end of shoot extension). The presence of honeydew significantly increased the growth of bacteria, yeast and filamentous fungi on the surface of needles and there was a pronounced seasonal trend, with the highest abundance in midsummer correlating with the period of peak aphid abundance. Taken together, these findings indicate that aphids have an influence on microbial ecology in the phyllosphere of trees. The implication of our study, from interactions at the population level to effects and potential consequences for C and N fluxes at the level of forest ecosystems, is discussed.     

The role of refuge areas in the phenology of Rhopalosiphum padi in low rainfall cropping areas of South Australia

Extensive surveys of possible aphid habitats in South Australia indicated that irrigated perennial grass pastures in the Mount Lofty Ranges and Lower Murray Valley were summer refuges for Rhopalosiphum padi (L.) (Hemiptera: Aphididae). Large numbers of aphids build up in these pastures each year during autumn (April and May) with numbers peaking in May. The size of the May peak was related to the number of aphids surviving the summer. The proportions of alates were highest in May and August/September. Both peaks coincided with a photoperiod of between 11.2 and 11.5 h, and partial correlations suggested that aphid density, photoperiod and temperature were all significant determinants of alate production.     

Populations of root-feeding aphids in the Liphook forest fumigation experiment

Three species of root aphid (Pachypappa vesicalis Koch, Pachypappa tremulae L. and Prociphilus (Stagona) xylostei De Geer) were found feeding on Norway spruce [Picea abies (L.) Karsten] that were exposed to elevated concentrations of O₃, and/or SO₂ in a field manipulation experiment at Liphook, England. The experiment exposed conifer trees to controlled concentrations of these gases between 1987 and 1990. Root-aphid populations feeding on Norway spruce were monitored in autumn 1990, the third year of fumigation and in autumn 1991, 10 months after fumigation had ceased. In 1990 the numbers of aphids in the high-SO₂, high-SO₂+ O₃ and O₃ treatments were significantly less than in ambient plots. Although after fumigation had ceased in 1991 the ambient plots still had higher numbers than the treatment plots, most of these differences were no longer statistically significant. In all plots sampled, total numbers of root aphids were higher in 1991 than in 1990. These results indicate that the numbers of root aphids in the experiment were negatively related to pollution levels. This relationship is the same as that shown in controlled laboratory experiments and is the converse of that generally reported in experiments using shoot-feeding aphids.     

Acclimatisation in the green spruce aphid, Elatobium abietinum

Thin layer chromatography separation of 80% ethanol extracts of adult Elatobium abietinum revealed the presence of the polyhydric alcohol mannitol in aphids overwintering outdoors but not in aphids kept permanently indoors at 15°C. After 3 days at 15°C no traces of mannitol were left in overwintering aphids. Mean freezing temperatures of outdoor, unfed instar I nymphs were about 4°C lower than those of unfed instar I nymphs produced at 15°C. Mean freezing temperatures of overwintering adults were considerably higher than those of unfed instar I nymphs and showed no changes associated with time at 15°C following transference indoors. Similarly, mean freezing temperatures of Sitka spruce needles transferred to 15°C did not change. It was concluded that, although freezing was mainly avoided by supercooling, the presence of mannitol lowered the true freezing temperature of aphid haemolymph and, consequently, the actual freezing temperatures of nymphs produced under cold conditions. However, the considerable increase in freezing point temperatures caused by imbibition of plant sap masked these acclimatisation changes in feeding aphids.     

Influence of sticky trap color and height above ground on capture of alate Elatobium abietinum (Hemiptera: Aphididae) in Sitka spruce plantations

A series of field trials were used to assess the practicality of using sticky traps to monitor populations of green spruce aphid, Elatobium abietinum (Walker), in plantations of Sitka spruce. The highest numbers of alate E. abietinum were caught on sticky traps placed in the upper third of the live canopy at 9-17 m above the ground, whereas low numbers of aphids were caught just below the live canopy or at 2 m above the ground. Trials in 2005 with sticky traps of different colors showed that significantly more alate E. abietinum were caught on yellow, red, and green sticky traps than on white, blue, and black traps. A repeat trial in 2007 resulted in significantly more alate aphids being caught on red sticky traps than on traps of any other color except for green. Attraction to red is unusual among aphids, as aphids are thought not to possess a red-sensitive photoreceptor. The attraction of E. abietinum to red-colored sticky traps suggests that conifer-feeding aphids might have a fundamentally different color response compared with aphids that live on cereals, grasses, or herbaceous plants. Alternatively, the attraction to red might be a physiological artifact related to the presence of red-screening pigments in the aphid's compound eye.     

区域性农田景观格局对棉蚜种群数量的生态学效应

农田景观格局的变化显著影响害虫的发生和危害,不同景观格局会对害虫的种群数量产生不同程度的影响,因而明确农田景观格局对害虫的生态学效应是控制害虫的重要前提之一。以山东省的棉花种植区为研究区域,选取14个典型的尽量临近不同土地覆盖类型的棉花生产县,通过卫星遥感影像和土地覆盖分类数据综合分析获得取样县/区的景观因子指数,并系统调查对应县/区的棉蚜种群数量。省级范围的大空间尺度下分析景观组成、景观构成和景观结构等多因子分别与棉田中苗蚜和伏蚜种群的相关性。研究结果表明棉蚜的种群数量与景观格局有密切的关系,且棉蚜发生的两个时期苗蚜和伏蚜对景观因子的响应特征并不完全一致。苗蚜的种群数量与景观总面积、耕地的分形指数、县域范围的蔓延度和县域范围的回旋半径等呈显著正相关,与Simpson多样性指标呈显著负相关;伏蚜的种群数量与斑块丰富密度、居住工业交通的蔓延度等呈显著正相关。总之,苗蚜和伏蚜对景观的蔓延度(形)响应基本上是一致的,景观的破碎化程度越小,伏蚜和苗蚜发生越重。而苗蚜和伏蚜对景观多样性(质)的响应不一致,景观多样性高的农田景观不利于苗蚜的发生,对伏蚜的影响不显著;而丰富度密度有助于伏蚜的发生,却对苗蚜没有显著影响。这一结果显示了农业害虫的不同发生时期对农田景观格局响应的复杂性。     

棉花型和黄瓜型棉蚜对寄主植物的适合度

采用寄主转换建立生命表及触角电位 (EAG)方法比较了棉蚜两寄主专化型 (棉花型和黄瓜型 )对不同夏季作物的适合度。结果表明 ,棉花型蚜转到黄瓜上及黄瓜型蚜转接到棉花上均不能正常产仔繁殖和建立种群。两种蚜型均不能在茄子和苋菜上建立种群。黄瓜型蚜能在豇豆上建立种群 ,但棉花型蚜不能 ,表现出两种寄主型棉蚜对夏寄主利用上存在显著差异。受棉蚜为害 12～ 36 h后的黄瓜或棉花植株仍不适合于棉花型或黄瓜型蚜 ,表现出黄瓜型蚜不能在被棉花型蚜为害过的棉株上正常存活和繁殖 ,棉花型蚜也不能在被黄瓜型蚜为害过的黄瓜苗上存活和繁殖。两种寄主型蚜对不同寄主叶片丙酮提取物的触角电位表明 ,黄瓜型蚜对棉花、哈密瓜和马铃薯叶片提取物的反应显著强于棉花型蚜 ,而对黄瓜和甜瓜叶片提取物的反应上两种蚜型差异不显著。文中同时对棉花型和黄瓜型棉蚜产生的生态机制进行了讨论。